Forest edge and diversity: carabids along forest-grassland transects

Diversity relationships of carabids in forest edges and the neighbouring forest interior and the surrounding grassland are studied. Samples were taken along three replicated forest-grassland transects using pitfall traps in the Aggtelek National Park in Hungary during 2 years. The study revealed significant edge effect on the carabids. The Shannon diversity of carabids were significantly higher in the forest edge and the grassland than in the forest interior. Carabids of the forest interior, forest edge and grassland can be separated from each other by ordinations, both on the species composition and abundance, suggesting that all three habitats have a distinct species assemblage. Moreover, indicator species analysis detected significant edge associated species; based on the specificity and fidelity of the carabids we have distinguished five groups of species: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analyses showed that relative air moisture, temperature of the ground and cover of the herbs are the most important factors determining the diversity of carabids along the transects. Our results show that the high diversity of carabids in forest edges is due to the edge-associated species and the presence of species characteristic to the adjacent habitats. The significance of forest edges in nature conservation, serving as a source habitat for dispersal processes, contributing to the recolonization of carabids after habitat destruction or other disturbance is emphasized.     

In situ extinction of carabid beetles and community changes in a protected suburban forest during the past century: the “Bosco Farneto” near Trieste (Italy)

The carabid beetle species assemblages (14 sample sites) of a 238 ha urban oak forest in Trieste, Italy, studied in 1983–84 with pitfall traps, were compared with an historic list of 57 species hand collected by entomologists in the same forest before it was logged in 1944.Trap data have been improved by hand collections to get a species list as complete as the historic one. Multivariate analysis was used to group the sites (14 plus the historic list) into three assemblages of brooks, clearings and forests. After the Second World War logging, the secondary ecological succession resulted in a lower species number, with a trend to a new equilibrium. Some important forest specialists, such as Laemostenus venustus, have been lost. Human pressure and recolonisation by carabids are still occurring. The in situ extinction of ground beetles near the end of the last century reached values between 57 and 64% of the species historically recorded. Consequently, urgent restoration measures are required for waterside habitats, forest and land management, to maintain small open areas or clearings. The study of carabid species assemblages and habitat affinities seems a useful tool for adaptive management of forests affected by human activities, because changes in carabid species number and type can be easily related to human disturbance.     

小兴安岭阔叶红松林地表甲虫Beta多样性

Beta多样性用来衡量集群内物种组成的变异性,可以被分解为空间物种转换和物种集群镶嵌两个组分,是揭示群落构建机制的重要基础。目前开展了较多的地上生态系统beta多样性研究,然而地下生态系统beta多样性进展缓慢。以小兴安岭凉水和丰林自然保护区为研究地区,于2015年8、10月采用陷阱法对阔叶红松林进行调查,揭示地表甲虫(步甲科、隐翅虫科、葬甲科)的beta多样性。结果表明:(1)凉水共发现39种、856只地表甲虫,丰林共发现43种、1182只地表甲虫。8月凉水明显具有较高的全部甲虫(三个科的总和)物种多样性和丰富度,10月正好相反。(2)凉水和丰林之间地表甲虫beta多样性的差异仅发现于8月的步甲科和葬甲科之间。(3)凉水和丰林地表甲虫的beta多样性主要由空间物种转换组成,物种集群镶嵌对beta多样性的贡献很小,说明地表甲虫物种组成变异主要由本地物种之间较高的转换引起。研究表明小兴安岭阔叶红松林地表甲虫的beta多样性主要由空间物种转换组成,在揭示群落构建机制过程中,其内部物种交换和环境调控不容忽视。     

Forest edge contrasts have a predictable effect on the spatial distribution of carabid beetles in urban forests

Fragmented urban forest remnants are characterised by sharp edges and are bordered by various land-use types, which may have a considerable effect on the fauna and flora at forest edges, and into forest interiors. To investigate the effects of differentially contrasting edges (low vs. intermediate vs. high) on carabid beetle assemblages in urban boreal forests, we placed pitfall traps along a gradient from 6?m into three matrix types (secondary forest vs. grassland vs. asphalt) up to 60?m into urban forest patches in the cities of Vantaa and Helsinki, southern Finland. Individual species and carabid beetle assemblages were strongly affected by edge contrasts and distance from the forest edge. The strongest effect on individual species was caused by high contrasting edges: generalist and open-habitat species were favoured or not affected while forest specialists were affected negatively. Effects of the abundances of potential prey and competitors on the carabid beetles were also evaluated. Forest and moisture-associated carabid species were negatively to neutrally associated with springtail abundances while generalist and open habitat, and dryness associated species were more positively related to springtail abundances (a potential food source). In terms of potential competitors, forest and moisture-associated carabid species were negatively and/or neutrally affected by ant and wood ant numbers, while generalist and open-habitat species were neutrally to positively associated with these taxa. It appears that carabid beetle habitat associations are more important in the responses of these beetles across edges of different contrast than are the prey and competitor numbers collected there. We recommend the creation of “soft” or low-contrast urban edges if the aim of urban management is to protect forest carabids in cities.     

The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient

Heterogeneity in species assemblages of forest-floor arthropods — carabid beetles, ants and spiders — within and between different forest age classes was studied in the southern Finnish taiga. The importance of processes operating on the local scale (within the movement radius of the species) vs on the regional scale (among forest stands) in determining the observed variation was assessed. Four data sets with different spatial resolutions in mesic forests in the same general study area were used. The material consists of 18 283 carabids of 51 species, 48 769 spiders of 212 species, and 126 718 worker ants of 23 species. Analyses of abundance variation and species complementarity among successional stages revealed that in all the three taxa species occurring in the mature forest were prevalent in the younger successional stages as well, constituting more than half of the catch in any age class. A great majority of carabid and spider species were widely distributed across the forest age classes, whereas ants include a higher proportion of species with a narrower amplitude across the succession gradient. Comparisons of similarity between samples at increasing distance from one another on the local scale within forest stands (a few tens of meters to a few hundreds of meters) showed a quite consistent pattern in carabids and spiders: there was more variation between sampling sites in young successional forests than in the mature forest. Furthermore, only in the mature forest a slight, albeit statistically not significant, negative relationship between similarity of samples and distance between sampling sites was detected. In carabids and spiders, comparisons between samples located at a distance of 10–15 m from each other showed considerable heterogeneity, the mean percentage similarity being c. 0.6 (in ants c. 0.8). On the regional scale, systematic variation between young and mature forest stands is a major element increasing the total diversity (species turnover c. 50% in carabids and spiders; compositional similarity c. 0.3–0.4 in carabids, 0.2–0.3 in spiders), but variation within forest stands on a spatial scale of 10–15 meters is another important component in the total heterogeneity. The results suggest that regional abundance variation is a primary factor influencing the composition of local assemblages; a set of hypotheses elaborating this conclusion is formulated. The result implies that maintenance of habitat heterogeneity on a small scale (10–15 m) is needed to preserve biodiversity in managed forests.     

Arthropod Diversity in Lama Forest Reserve (South Benin), a Mosaic of Natural, Degraded and Plantation Forests

Arthropod assemblages were examined in Lama forest reserve, a protected area situated in the Dahomey gap, southern Benin, composed of plantations, degraded forest and remnants of natural forest. The objectives were to compare assemblages in relation to forest type and use, to elucidate the value of forest plantations for biodiversity conservation and to identify indicator species for specific forest habitats. Arthropods were collected over an 11-month period, using standardized sets of traps (pitfall, emergence, Malaise and flight intercept traps). Nine different habitats were studied, including natural and degraded forest, forest plantations (Tectona grandis and Senna siamea) of different age, and isolated forest fragments. Our analysis focused on detritivorous and xylophagous arthropods but also included ground beetles and heteropterans, totalling 393 species. We found no differences in species richness among natural and degraded forest habitats in the centre of the reserve (Noyau central). Outside of the Noyau central, species richness was highest in old teak plantations and isolated forest fragments and lowest in young teak and fuelwood plantations. Detrended correspondence analysis (DCA) separated three main groups: (1) natural forest, (2) degraded forest and young plantations, and (3) old plantations and isolated forest fragments. Multiple regression of DCA scores of the first two axes on environmental variables identified one natural and three disturbance-related predictors of arthropod assemblages in Lama forest: soil type (texture), canopy height, naturalness (proportion of Guineo-Congolian plant species) and understorey vegetation cover. We identified 15 indicator species for six different forest habitats. The highest numbers were found in abandoned settlements and old teak plantations. β-diversity was similar among the three DCA ordination groups (degraded forest excluded). Values for β-diversity were relatively high, suggesting that all major forest habitats contribute significantly to regional species pools and should therefore be protected. To enhance arthropod diversity, we propose that management practices in Lama forest should aim to encourage the development of species-rich understorey vegetation of the Guineo-Congolian phytogeographical region.     

Response of ground and rove beetles (Coleoptera: Carabidae,Staphylinidae) to operational oil sands mine reclamation in northeastern Alberta,a case study

Species loss caused by anthropogenic disturbance threatens forest ecosystems globally. Until 50 years ago, the major sources of boreal forest disturbance in western Canada were a combination of forest wild fire events, pest insect outbreaks, and forest timber harvesting. However, in the 1960s, when the oil boom started in Alberta, oil and gas development along with oil sands mining quickly became another major forest disturbance agent. In this case study we report the effects of operational oil sands mine reclamation on terrestrial arthropod communities and compare them with nearby burned and mature forest sites as a way to provide a benchmark from which to understand the long-term trajectory of recovery for these groups. During the summer of 2016 over 6700 epigaeic beetles were collected using pitfall traps. A total of 43 species of ground beetles and 118 species of rove beetles were collected. Epigaeic beetle assemblages differed between the reclaimed, burned, and mature forest sites. Partitioning of beta diversity in the reclaimed, burned areas and mature forests indicated that species turnover formed the largest component of diversity. Species richness patterns were similar among sites; however, cluster analysis indicated that epigaeic beetle assemblages were only 20% similar between the reclaimed and natural sites. Although ground beetles of the reclaimed area showed positive spatial autocorrelation among treatments, both ground and rove beetles showed responses to the reclamation treatments. The reclaimed areas were dominated by small- to medium-sized open-habitat eurytopic species, whereas the fire and mature forest sites were dominated by larger forest species. The reclaimed area of this case study constitutes a novel, reconstructed ecosystem that is clearly not equivalent in species assemblage to burnt stands of similar age or to mature forest stands.     

Are streamside buffers edge-affected habitat for ground-dwelling forest beetle assemblages?

Reservation of forest in riparian buffers is common practice in commercial forestry areas worldwide, potentially providing valuable habitat for biodiversity dependent on mature forest. However, the habitat value of narrow reserve corridors can be compromised by edge effects. We investigated the habitat value of streamside buffers in wet eucalypt forest for ground-dwelling beetles in Tasmania, Australia. Beetles were collected with pitfall traps in five replicates of four habitats: unlogged corridors of mature forest in streamside reserves (buffers) with clearfelling-derived logging regeneration either side; continuous mature upslope forest; continuous mature riparian forest; and <20-year-old upslope clearfelling-derived logging regeneration. Streamside reserve widths on each side of the stream were on average 40 ± 6 m (±95% CI) from reserve edge to stream. Beetle assemblages in logging regeneration differed substantially from those in the unlogged habitats, including the streamside reserves. Streamside reserve assemblages nevertheless differed from those of the continuous unlogged areas. Assemblage composition in streamside reserves was most similar to that in continuous mature riparian forest, although beetle diversity was higher in the reserves. Our results suggest that although streamside reserves provide habitat for the majority of commonly collected beetle species occurring in continuous mature forest, wider reserve corridors in the wet eucalypt forests of Tasmania may be required to provide habitat that is not edge-affected.     

Carabid beetles (Coleoptera: Carabidae) at forest-farmland edges in southern Finland

We studied carabid beetle abundance at eight forest-farmland edges using pitfall traps across 60-m gradients (30 m into the forest, 30 m into the adjacent farmland) in southern Finland in May–August 2001. Carabid assemblages changed gradually across the studied gradients, the most drastic changes occurring right at the edge (5 m). Forest-associated carabids were often caught in farmland habitat within 20–30 m from the edges, and open-habitat carabids were also caught in the forest patches. However, these two groups responded to the edge in slightly different ways. Forest carabids were abundant all across the gradient from forest interior to the edge ( 80 m 9), while open-habitat carabids showed a drastic abundance decrease toward the forest, 5–10 m before the edge the abundance-change slope across the edge: gradient was steeper for open-habitat than for forest carabids. Wing-dimorphic and long-winged carabids increased more steeply from forest to farmland, compared to short-winged carabids. Moreover, carabids associated with dry and moist habitat showed indications of stronger response to the edge than did eurytopic species. The pair-wise comparisons between predatory/mixed-diet carabids and seed-eaters, spring and autumn breeders, and day- and night-active species did not indicate edge-response differences.     

Dead trees left in clear-cuts benefit saproxylic Coleoptera adapted to natural disturbances in boreal forest

Forest management alters the pattern of forest dynamics from that in natural conditions in the boreal region. In order to examine how certain forestry measures matching natural dynamics affect forest insects, we compared assemblages of saproxylic Coleoptera on dead, standing birch trunks left behind in eight clear-cut areas with corresponding assemblages in seven mature forests in southern and eastern Finland. We used trunk-window traps for sampling. Distinct beetle assemblages were associated with the different habitats. Median numbers of species or specimens caught did not differ between closed forests and clear-cuts, but individual beetle species occurred unevenly among the habitats. Several beetle species associated with open forest habitat, e.g. burned forests or storm-damage areas, including species regarded as threatened in Finland, were found almost exclusively, in clear-cuts. Correspondingly, a number of beetle species occurring frequently in closed forests were not found in clear-cuts. We conclude that dead trunks left in the clear-cut areas may host not only generalist saproxylic species but also many beetle species specialized to warm, sun-exposed environments, and such species may not be able to survive in closed forests. Management measures matching suppressed natural disturbances are found useful in preserving diversity in managed forests.     

Oil pipeline corridor through an intact forest alters ground beetle (Coleoptera: Carabidae) assemblages in southeastern Ohio

Litter-dwelling ground beetle (Coleoptera: Carabidae) assemblages were monitored 1 yr after the construction of a corridor for installation of an oil pipeline along a xeric ridge-top forest in southeastern Ohio. After the creation of the corridor, three distinct habitats were evident in these sites: open corridor, ecotone areas around the corridor, and undisturbed forest interior. Carabidae were collected using directional pitfall traps that were placed parallel and perpendicular to the corridor in each of the three habitats. Results indicate that more carabids were present in the ecotone than in the other two habitats. Carabid diversity as estimated by rarefaction was highest in the corridor followed by ecotone and forest interior. Generalist and forest specialists such as Synuchus impunctatus (Say), Carabus goryi Dejean, and Pterostichus trinarius (Casey) were present in greater numbers in the forest interior and ecotone assemblages. In contrast, open-habitat specialists such as Harpalus pensylvanicus (DeGeer) and Selenophorus opalinus (LeConte) were present in greater numbers in the corridor assemblages. Carabid assemblages of the corridor were distinct from those of the ecotone and forest interior, whereas the latter two habitats had very similar assemblages. The successional pathway of the corridor carabid assemblage will therefore be likely different from that of the forest interior and ecotone. Overall, results indicate that construction of the oil pipeline corridor had significant short-term effects on the carabid numbers, diversity, and species composition because of ensuing habitat changes and fragmentation of the forest.     

Measuring the efficiency of fen restoration on carabid beetles and vascular plants: a case study from north‐eastern Germany

The aim of the study was to assess the effects of fen rewetting on carabid beetle and vascular plant assemblages within riverine fens along the river Peene in north‐eastern Germany. Drained (silage grassland), rewetted (restored formerly drained silage grassland), and near‐natural (fairly pristine) stands were compared. Eighty‐four beetle species (7,267 individuals) and 135 plant species were recorded. The richness of vascular plant species and the number of endangered species were highest on near‐natural fens. Fourteen years of rewetting did not increase plant species numbers compared with drained fens. For carabid beetles, however, species richness and the number of stenotopic species were highest on rewetted fens. Rewetting caused the replacement of generalist carabids by wetland specialists, but did not provide suitable habitat for specialist fen carabids or for plant species of oligo‐ or mesotrophic fen communities. Therefore, raising the water table on fens with nutrient‐rich, degraded peat was not sufficient for restoring species assemblages of intact fens, although water level was the most important environmental factor separating species assemblages. Our study illustrated that insects and plants may respond differentially to restoration, stressing the need to consider different taxa when assessing the efficiency of fen restoration. Furthermore, species assemblages of intact fens could not be restored within 14 years, highlighting the importance of conserving pristine habitat.     

Responses of ground-active beetle assemblages to different styles of reforestation on cleared rainforest land

Despite increasing efforts to re-establish forest cover in landscapes that have been previously cleared, the relative ability of different styles of reforestation to contribute to conservation and support forest biota is poorly known, particularly for invertebrates. We investigated the use of different types of reforested habitat by ground-active rainforest beetle assemblages on land, which had been previously cleared of rainforest, in the tropics and subtropics of eastern Australia. Between five and ten replicate sites within each of five reforestation styles were selected in each region: un-managed regrowth, young mono-species timber plantations, young mixed-species timber plantations, ecological restoration plantings, and old mono-species timber plantations, together with reference sites in pasture and in intact rainforest. Ground-active beetles were sampled using pitfall traps, and assemblages were compared among site-types. In both regions, beetle assemblages in all styles of reforestation were intermediate in species composition between pasture and rainforest. The similarity of beetle assemblages to intact rainforest increased with the age and structural complexity of reforested sites. The most rainforest-like beetle assemblages were from older reforestation sites (38–70 year plantations in tropics, and 30–40 year regrowth in subtropics) and in younger (6–22 years) but floristically and structurally diverse ecological restoration plantings in both regions. Assemblages in younger (5–20 year) sites of regrowth, mono-species timber plantations, and mixed-species timber plantations were more similar to pasture than rainforest. We conclude that achieving high canopy cover and sufficient structural complexity are important factors associated with the restoration of rainforest-like beetle assemblages to reforested sites.     

Influence of sampling effort on saproxylic beetle diversity assessment: implications for insect monitoring studies in European temperate forests

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The effect of forest age and habitat structure on the ground‐dwelling ant assemblages of lowland poplar plantations

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Does selective logging change ground‐dwelling beetle assemblages in a subtropical broad‐leafed forest of China?

Selective logging with natural regeneration is advocated as a near‐to‐nature strategy and has been implemented in many forested systems during the last decades. However, the efficiency of such practices for the maintenance of forest species are poorly understood. We compared the species richness, abundance and composition of ground‐dwelling beetles between selectively logged and unlogged forests to evaluate the possible effects of selective logging in a subtropical broad‐leafed forest in southeastern China. Using pitfall traps, beetles were sampled in two naturally regenerating stands after clearcuts (ca. 50 years old, stem‐exclusion stage: selectively logged 20 years ago) and two mature stands (> 80 years old, understory re‐initiation stage: selectively logged 50 years ago) during 2009 and 2010. Overall, selective logging had no significant effects on total beetle richness and abundance, but saproxylic species group and some abundant forest species significantly decreased in abundance in selectively logged plots compared with unlogged plots in mature stands. Beetle assemblages showed significant differences between selectively logged and unlogged plots in mature stands. Some environmental characteristics associated with selective logging (e.g., logging strategy, stand age, and cover of shrub and moss layers) were the most important variables explaining beetle assemblage structure. Our results conclude that selective logging has no significant impacts on overall richness and abundance of ground‐dwelling beetles. However, the negative effects of selective logging on saproxylic species group and some unlogged forest specialists highlight the need for large intact forested areas for sustaining the existence of forest specialist beetles.     

Patch size determines the strength of edge effects on carabid beetle assemblages in urban remnant forests

Habitat fragmentation due to urbanization is increasing rapidly worldwide. Although patch area and edge effect are both important determinants of species diversity and the number of individuals in fragmented landscapes, studies that tested interaction between two effects were limited. Here we examined the interaction between area and edge effects on species richness and the number of individuals of carabids in highly fragmented forests in Tokyo, central Japan. We surveyed carabids in each of 26 forest patches (1.1–121.6 ha) using pitfall traps set in both edge and interior zones. First, we correlated the edge-to-interior differences of both species richness and the number of individuals with patch area. Second, we examined the interaction between patch area and distance to the edge on species richness and the number of individuals using generalized linear models. We found a significant decrease in carabid species richness and the number of individuals in edge zones. The edge-to-interior differences in both species richness and the number of individuals were positively correlated with patch area. Model selection revealed the evident interaction effects between patch area and distance to the edge: higher number of individuals was predicted in only large interior zones. Our results indicated that carabid beetle assemblages were influenced by the interaction between area and edge effects. Thus, in urban areas where small forest remnants dominate, circularizing the shape of the forest patches to maximize the core areas may be the most feasible and realistic means to preserve biodiversity.     

Ground beetle diversity in ancient woodland remnants in north-western Germany (Coleoptera, Carabidae)

We investigated ground beetle communities (Coleoptera, Carabidae) in ancient woodland remnants in north-western Lower Saxony, Germany. A total of 90 pitfall traps was exposed in a stratified design in 10 stands of mature oak–beech and oak–hornbeam forests in the year 2003. Overall, 47 species (10,676 individuals) were recorded. Among these were the two relict species Carabus glabratus and Abax parallelus, and 14 further eurytopic forest species. Eleven species exhibited a high frequency and were found in all of the ten stands. Multiple linear regressions showed several significant relationships at two scales for species richness of different groups of carabids and for several of the measured environmental factors. Forest area, litter depth, amount of dead wood, distance to forest edge, and soil moisture were found to be key factors determining species richness. Furthermore, recent disturbance by logging reduced the number of forest species. According to direct gradient analyses soil moisture and litter depth have greatest influence on species communities of both, forest species and widespread species. Habitat suitability models for the two recorded relict species, A. paralellus and C. glabratus, were developed using logistic regression. The presence of A. parallelus in the mature ancient woodland remnants depends mainly on higher values of soil moisture, whereas for C. glabratus none of the measured environmental variables appeared to be key factors. Implications for the conservation of carabid assemblages in mature ancient woodlands include the advice to spread out logging over long periods of time and over various woodlands in order to keep the stand disturbance at a long-term low level. Variation in logging practices may help to conserve diverse structures. Afforestation with non-native tree species should be avoided in the managed ancient woodlands. Finally, especially the preservation of a high soil moisture seems to be important to conserve typical carabid communities.     

Sampling indigenous ground‐living beetles in a stand of non‐native tree privet (Ligustrum lucidum) in New Zealand raises new management questions

Small urban forest reserves in New Zealand have been shown to have value in conserving indigenous beetle diversity. However there is little information available on the ability of non‐native vegetation areas such as tree privet to support indigenous beetle assemblages. To investigate this for one site, ground‐living beetles were collected using pitfall traps over a year at a small urban forest of the invasive tree Ligustrum lucidum (tree privet) in Auckland, New Zealand. A total of 815 beetles were found, from 20 families and 42 relative taxonomic units. Using monthly data, there was no correlation between soil moisture and diversity index (P = 0.805) or species richness (P = 0.375). These results raise the question of whether urban patches of non‐native tree privet may have potential as reservoirs of beetle diversity, if only until they are replaced with native vegetation.